4
DEC. 7, 1912.] FUNCTION OF CAECUiVE AND APPENDIX. TICB '599 lis the consequence, and not the cause, of the arrest of evolution. Tumour tissues are imperfect- and remain ,imperfect, and work cannot be expected of such. Their differentiation has been arrested before they were capable of work. Tumour tissues never work because, with the rarest possible exceptions, they never reach the perfection of maturity. The exception proves the rule; a few mammary adenomata attain such a high degree of p3r- fection that secretion has been known to take place in certain portions of them. The vast majorityfalls far short of this. THE ARREST OF DEVELOPMENT OF ORGANS. If it is asked what could interfere with the power of hereditary transmission to such an extent as to cause the arrest of the differentiation of the tissues in one part of the body and the formation of one kind of tumour, or the arrest of their involution in another and the formation of another kind, I can only point to the analogy of the arrest of development of organs, and say that I can see no reason why such a power may not fail from mere exhaustion, as for instance in old age, either of organs or of individuals, when it is notorious that the liability to tumour growtl becomes greater and greater, or from excessive action, as when an enormous number of young cells is produced with great rapidity in response to some persistent clhronic irritation. Whether the action, or want of action, of any organ in the body could have a similar influence over the power of hereditary transmission and bring its work to a sudden stop it is impossible to say. There is one struc- ture in the body at least that appears to be capable of abrogating the laws that control the growth of tissues and of enabling them to increase in size without limit; but we know too little of these things as yet. The study of physiology is of yesterday. At our best we have only a very general acquaintance with some of the most con- spicuous individual actions of what may be called the gross mechanical apparatus of the body. We know next to nothing of the essential organs of life, those that really matter where growth and development are concerned. We are only just beginning to grasp the fact that many absolutely insignificant-looking structures, some of them almost microscopic in size, possess the most extraordinary power over growth, and even over life itself,; and we are only just beginning to realize that no organ or tissue in the body exists for itself alone, but only in relation with every other organ and tissue. It may be tllat when we linow more about these things some structure will be found which has some control over lhereditary trans- mission and the development of tissues as distinguished from their growth. Until then-and it must be admitted the prospect is not a good one-all our hopes of finding suclh a thing as a cure for the growth of tumours (for amputation, though it may free the patient from the con- sequences, can in no sense be regarded as a cure) must rest either upon finding some remedy which, like Coley's fluid, can directly kill the tumour cells without endanger- ing too greatly the vitality of the normal tissues, or upon stimulating to increased energy the normal tissues that surround the tumour until they are able to treat those degenerate cells that have failed to attain their full development and have continued to grow at the expense of the organism without being of any service to it, in the same way that the workers treat the drones. DR. EDGAR BATES has examined all the children in the Dee and Washington schools of Ogden, Utah; 890 pupils were tested with the illiterate card, and 184 were found to be suffering from eyestrain. Of these 134, or 73 per cent., had headaches; 114 complained of symptoms originating in close work, and 57 of them stated that words became blurred; 34 had blepharitis. A gradual increase of cases was noted from the kindergarten up to the highest forms. Many of the pupils stated that their symT)toms were first noted after an attack of measles or scarlet fever. Some of the children who complained of eyestrain were wearing glasses, but they had been fitted by an optician without the use of a mydriatic, which Dr. Bates considers futile; most ophthalmologists will subscribe to this opinion. In reading this paper, which appears in Ophthalmology for January, 1912, it is necessary to remember that eyestrain is far more frequent in America than it is here, and also that leading questions as regards headache were put to the children. THE FUNCTIONAL NATURE OF THE CAECUM AND APPENDIX.* BY ARTHUR KEITH, M.D., CONSERVATOR OF MUSEUM, ROYAL COLLEGE OF SURGEONS OP ENGLAND. EVERY year the opinion gains ground that the great bowel, from appendix to rectum, has become, so far as muan is concerned, a useless and dangerous structure. Exactly ten years ago (October, 1902) Dr. Barclay Smith1 of Cambridge gave a clear expression of this newr conception. " If there is any truth," he wrote, "in the suggestions offered in this paper, they have an important practical application as regards the large intestine of man. From the nature of his diet a reliance on extrinsic digestive aid as is furnished by bacteria is no longer a physiological necessity. The statement is perhaps a bold one, but I am convinced that the large intestine is practically a useless encumbrance to him."' In the following year (1903) appeared Metchnikoff's famous book on The Natutre of Man.2 In that work (p. 69) the new conception receives a more decisive statement: It is no longer rash to sav that not onlv the rudimentary appendix and the caecum, but the whole of the large intestine are superfluous, and that their removal would be attended with happy results. Before either of these statements had been made, Mr. Arbuthnot Lane3 had reached the conclusion that the lhuman caecum and ascending colon served, in a certain class of cases described by him, as a "c esspool," and put his new conception into practice either by excluding the great intestine from the digestive tract by " short-circuiting," or, at a later date, by its, complete excision. The result of his operations shows (1) that life is possible without a great intestine; (2) that in certain cases the conditions of life are improved. It is very apparent that Mr. Lane's pioneer operative measures. are finding every year an increasingly wide application, and that the views of Metchnikoff and of Barclay Smith are gaining in acceptation amongst medical men. THEORIES REGARDING THE USES OF THE CAECUM AND COLON. If these statements are well founded-if the great bowel is as useless and injurious as is said then medical men are face to face with a condition which threatens the health and survival of modern civilized'races. Before assigning so large a part of man's digestive tract to the list of his useless structures, it will be well to inquire what we know regarding the function of the great bowel as a whole, and the uses of its parts in the various members of the animal kingdom. Such an inquiry brings home to us that our knowledge of the function and sig- nificance of the great bowel is vague and unsatisfactory. We do not know, in any animal whatsoever, the exact function or functions of the great bowel; so far we have only guessed at them. The subject is one which engaged the attention of John Hunter; in his collection there are many specimens which illustrate the structure of the various parts of the great bowel in all classes of vertebrates, but unfortunately he left only the briefest record of the conclusions he had formed regarding their uses. "If we could understand," he wrote, "the use of the ileo-eaccal valve, then we should understand the uses of the caecum."4 He observed that there was a material difference between the contents of th3- ileum and of the caecum; the contents of the caecum and colon underwent putrefactive changes, and in his opinion the ileo-caecal value was to prevent the putrid contents of the caecum from regurgitatinig within the ileum. In a case of gun-shot wound of the abdomen he observed that the colic contents lhad passed upwards in the ileum, a result which he supposed might be due either to a lesion of the nerve supply of the bowel or to a reversed action ot the intestine. In the light of our modern knowledge it is interesting to note he lhad observed that the colic contents were not putrid at birth nor did the caecum of the newly-born conltain gas. He was also famliliar with the fact that the caecuim was small, the colon short, and '+ Abostract of a demonlstration at the College.-

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Page 1: THE FUNCTIONAL NATURE OF THE CAECUM AND APPENDIX.*

DEC. 7, 1912.] FUNCTION OF CAECUiVE AND APPENDIX. TICB '599

lis the consequence, and not the cause, of the arrest ofevolution. Tumour tissues are imperfect- and remain,imperfect, and work cannot be expected of such. Theirdifferentiation has been arrested before they were capableof work. Tumour tissues never work because, with therarest possible exceptions, they never reach the perfectionof maturity. The exception proves the rule; a fewmammary adenomata attain such a high degree of p3r-fection that secretion has been known to take place incertain portions of them. The vast majorityfalls far shortof this.

THE ARREST OF DEVELOPMENT OF ORGANS.If it is asked what could interfere with the power of

hereditary transmission to such an extent as to cause thearrest of the differentiation of the tissues in one part ofthe body and the formation of one kind of tumour, or thearrest of their involution in another and the formation ofanother kind, I can only point to the analogy of the arrestof development of organs, and say that I can see no reasonwhy such a power may not fail from mere exhaustion, asfor instance in old age, either of organs or of individuals,when it is notorious that the liability to tumour growtlbecomes greater and greater, or from excessive action, aswhen an enormous number of young cells is producedwith great rapidity in response to some persistent clhronicirritation. Whether the action, or want of action, of anyorgan in the body could have a similar influence over thepower of hereditary transmission and bring its work toa sudden stop it is impossible to say. There is one struc-ture in the body at least that appears to be capable ofabrogating the laws that control the growth of tissuesand of enabling them to increase in size without limit;but we know too little of these things as yet. The studyof physiology is of yesterday. At our best we have onlya very general acquaintance with some of the most con-spicuous individual actions of what may be called thegross mechanical apparatus of the body. We know nextto nothing of the essential organs of life, those that reallymatter where growth and development are concerned.We are only just beginning to grasp the fact that manyabsolutely insignificant-looking structures, some of themalmost microscopic in size, possess the most extraordinarypower over growth, and even over life itself,; and we areonly just beginning to realize that no organ or tissue inthe body exists for itself alone, but only in relation withevery other organ and tissue. It may be tllat when welinow more about these things some structure will befound which has some control over lhereditary trans-mission and the development of tissues as distinguishedfrom their growth. Until then-and it must be admittedthe prospect is not a good one-all our hopes of findingsuclh a thing as a cure for the growth of tumours (foramputation, though it may free the patient from the con-sequences, can in no sense be regarded as a cure) mustrest either upon finding some remedy which, like Coley'sfluid, can directly kill the tumour cells without endanger-ing too greatly the vitality of the normal tissues, or uponstimulating to increased energy the normal tissues thatsurround the tumour until they are able to treat thosedegenerate cells that have failed to attain their fulldevelopment and have continued to grow at the expenseof the organism without being of any service to it, in thesame way that the workers treat the drones.

DR. EDGAR BATES has examined all the children in theDee and Washington schools of Ogden, Utah; 890 pupilswere tested with the illiterate card, and 184 were found tobe suffering from eyestrain. Of these 134, or 73 per cent.,had headaches; 114 complained of symptoms originatingin close work, and 57 of them stated that words becameblurred; 34 had blepharitis. A gradual increase of caseswas noted from the kindergarten up to the highest forms.Many of the pupils stated that their symT)toms were firstnoted after an attack of measles or scarlet fever. Some ofthe children who complained of eyestrain were wearingglasses, but they had been fitted by an optician withoutthe use of a mydriatic, which Dr. Bates considers futile;most ophthalmologists will subscribe to this opinion. Inreading this paper, which appears in Ophthalmology forJanuary, 1912, it is necessary to remember that eyestrainis far more frequent in America than it is here, and alsothat leading questions as regards headache were put tothe children.

THE FUNCTIONAL NATURE OF THECAECUM AND APPENDIX.*

BY ARTHUR KEITH, M.D.,CONSERVATOR OF MUSEUM, ROYAL COLLEGE OF SURGEONS OP

ENGLAND.

EVERY year the opinion gains ground that the greatbowel, from appendix to rectum, has become, so far asmuan is concerned, a useless and dangerous structure.Exactly ten years ago (October, 1902) Dr. Barclay Smith1of Cambridge gave a clear expression of this newrconception." If there is any truth," he wrote, "in the suggestions offered

in this paper, they have an important practical application asregards the large intestine of man. From the nature of hisdiet a reliance on extrinsic digestive aid as is furnished bybacteria is no longer a physiological necessity. The statementis perhaps a bold one, but I am convinced that the largeintestine is practically a useless encumbrance to him."'In the following year (1903) appeared Metchnikoff's

famous book on The Natutre of Man.2 In that work(p. 69) the new conception receives a more decisivestatement:

It is no longer rash to sav that not onlv the rudimentaryappendix and the caecum, but the whole of the large intestineare superfluous, and that their removal would be attended withhappy results.

Before either of these statements had been made,Mr. Arbuthnot Lane3 had reached the conclusion thatthe lhuman caecum and ascending colon served, in acertain class of cases described by him, as a "c esspool,"and put his new conception into practice either byexcluding the great intestine from the digestive tract by" short-circuiting," or, at a later date, by its, completeexcision. The result of his operations shows (1) that lifeis possible without a great intestine; (2) that in certaincases the conditions of life are improved. It is veryapparent that Mr. Lane's pioneer operative measures. arefinding every year an increasingly wide application, andthat the views of Metchnikoff and of Barclay Smith aregaining in acceptation amongst medical men.

THEORIES REGARDING THE USES OF THE CAECUM ANDCOLON.

If these statements are well founded-if the greatbowel is as useless and injurious as is said then medicalmen are face to face with a condition which threatens thehealth and survival of modern civilized'races. Beforeassigning so large a part of man's digestive tract to thelist of his useless structures, it will be well to inquirewhat we know regarding the function of the great bowelas a whole, and the uses of its parts in the variousmembers of the animal kingdom. Such an inquiry bringshome to us that our knowledge of the function and sig-nificance of the great bowel is vague and unsatisfactory.We do not know, in any animal whatsoever, the exactfunction or functions of the great bowel; so far we haveonly guessed at them.The subject is one which engaged the attention of John

Hunter; in his collection there are many specimens whichillustrate the structure of the various parts of the greatbowel in all classes of vertebrates, but unfortunately heleft only the briefest record of the conclusions he hadformed regarding their uses. "If we could understand,"he wrote, "the use of the ileo-eaccal valve, then we shouldunderstand the uses of the caecum."4 He observed thatthere was a material difference between the contents ofth3- ileum and of the caecum; the contents of the caecumand colon underwent putrefactive changes, and in hisopinion the ileo-caecal value was to prevent the putridcontents of the caecum from regurgitatinig within the ileum.In a case of gun-shot wound of the abdomen he observedthat the colic contents lhad passed upwards in the ileum,a result which he supposed might be due either to a lesionof the nerve supply of the bowel or to a reversed action otthe intestine. In the light of our modern knowledge itis interesting to note he lhad observed that the coliccontents were not putrid at birth nor did the caecum ofthe newly-born conltain gas. He was also famliliar withthe fact that the caecuim was small, the colon short, and

'+Abostract of a demonlstration at the College.-

Page 2: THE FUNCTIONAL NATURE OF THE CAECUM AND APPENDIX.*

FUNCTION OF CAECUM AND APPENDIX.

the faeces of thin consistency in carnivora and alliedanimals, while the opposite was the case in vegetablefeeding animals. Further, he recognized that animalswith a perfect gastric digestion, such as ruminants, hadcomparatively small caeca, while in others, like the horse,with an imperfect gastric digestion, the caeca were large.He therefore recognized, a very important fact, that therecould be an interchange of function between stomach andcaecum. The task of describing and interpreting Hunter'sspecimens fell to Sir Richard Owen. The conclusion hedrew from an examination of the series of specimenswhich illustrate the anatomy of the ileo-caecal region ofthe bowel was:Comparative, anatomy concurs with the results of undesigned

experiments, as in. cases where artificial openings have beenestablished in the human intestinal canal, in showing that achange to gastric digestion is repeated upon the food in thecaecum; chemistry has also shown that the chyme here againbecomes acidified, after having been neutralized by bile in thesmall intestine.5

Both Hunter and Owen recognized that the process ofdigestion underwent a sudden and decided change at theileo-caecal orifice, but neither the one nor the othersuspected that- the digestive process was the result of aputrefactive action. That discovery, of course, belongs tomodern times.Reference has already been made to a paper1 which Dr.

Barclay Smith published ten years ago, one which has notreceived t h eattention 'its'importance de-s e r v e s. The SPHINCTERIC TRAC

theory fornu-.lated there hasthe merit ofseeking to ex-'plain the ana- "CAL COL

tomy of theileo-caecal re -gion. Histheory RETINAC:Bis based on thelustifiable as-sumption that, ILEU

all the digestive cLCL(eIchanges of foodwithin the greatbowel are due,not to any diges- ePN~tive secretion,but to the ac-tion of bacteria Fig. 1.-The human caecum laid open towhich find a show the ileo-caecal orifice, the retinacular

p bands, the caecal colon, and caeco-colicpae rman e n sphincteric tract. One-third natural size.abode in the-caecum and colon. He inferred-an inference whichhas been justified by the observations of Dr. Hertz -

that the antiperistaltic movements which Dr. Cannon7had seen to occur in the colon of the cat also occurredin the upper part of the human colon, and that thepurpose of the ileo-caecal valve was to prevent thecontents of the caecum and colon, which were undergoingbacterial digestion, from being forced by the antiperistalticmovements within the ileum, where a totally differentdigestion was in progress. So far as I know, this was thefirst satisfactory explanation offered for the presence of anileo-caecal valve. Dr. Barclay Smith looked on the rich dis-tribution of lymphoid tissue in the ileo-caecal region asstations furnished to protect the body from invasion by thebacterial hosts employed in the purposes of caecal digestion.He made the further assumption, which may or may notbe warrantable, that our modern diet needs no bacterialaction for its full digestion and assimilation, and thattherefore the great intestine is a useless and injuriousstructure in the economy of modern man. Excision, how-ever, is not the only remedy. There is an option; in placeof appealing to surgery to adapt our digestive tract to ourpresent dietary, it seems possible that we may dis-cover a diet which is suited to our present diges-tive tract. Before we can do that, we must knowsomething of the normal processes of digestion whichproceed within the 'great bowel, not 'only of manand of animals allied to him, but of vertebrate animalsgenerally.

ILEO-CAECAr, SPHINCTER.Johii Hunter, with his usual prescience, realized that

a knowledge of the ileo-caecal valve would provide a clueto the nature- of the caecum. In 1903, as a result of a.series of observatidns'on all classes of vertebrate animals,I' came to the conclusion that there was always afunctional muscular sphincter at the junction of thesmall intestine with thei great.8 It is present in theamphibian bowel, the reptilian, avian, and mammalian.When the great bowel' undergoes retrogression, as it hasdone ill bears and in their allies and in many forms ofinsectivora and edentates,9 the ileo-caecal sphincter is theonly visible structure-which remains to mark the junctionof th'e small intestine with the large. My observationsled me to regard the ileo-caecal sphincter as a constantstructure, its main function being to serve as a mechanismfor regulating the passage of the contents of the ileuminto the caecum, in much the same way as the pyloricsphincter regulates the passage of the food from thestomach into the duodenum.

Independently, Dr. T. R. Elliottl0 had observed the'ileo-colic sphincter in action, and placed our knowledge ofit on a much sounder basis. In May, 1904, he was able toannounce that stimulation of the sympathetic nerves of theinferior mesenteric plexus causes it to contract, and at thesame time the same stimulus brings about an inhibition ofthe neighbouring musculature of the ileum and colon;adrenalin, he found, had a similar effect. Dr. Barclay

Smith and he"concluded thatthe strength ofthe sphincterw a s commen-surate with thethickness of the

\o>< muscular coatof the caecum.

ILE--CAeC: ORIFtC0: a n d adjacentO*IFICE: < v ~p a rt of t heORIFIE _colon, and that

ECAL ORIFICE ILEUM) its purpose orfunction was toprevent thecaecal contentsb e i n g forcedbackwards into

-:CUMz _ the ileum underthe pressure

.1 - AP;PDIS caused by anti-p erista It i

Fig. 2.-The caecum of the rat (twice movements.natural size) laid open to show the same Sir W ill aparts as are seen in Fig. 1. M w 12 unaMacewen,l un-

aware of theobservations made by Dr. Elliott or by myself, announcedin the Huxley Lecture of 1904 that he had recognized thepresence of an ileo-caecal sphincter and studied its actionin a soldier who had a temporary window in the wallof his caecum. In this manner a most important stepforwards was made in our knowledge of the caecum, wecame to see that there is an elaborate neuro-muscularmechanism which regulates the passage of chyme intothe caecum and prevents the regurgitation of the contentsof the caecum into the ileum. The significance of theileo-caecal orifice and sphincter is clear if we admit thatthey mark a point in the alimentary tract where one formof digestion ends and another begins.

FUNCTIONAL DIVISIoNs OF THE CAECUM.In my preliminary studies 8 of the ileo-caecal region of

the bowel, I came to the conclusion that in the fullydeveloped mammalian caecum, such as that of the rabbilor rat, there are three parts which appear to bedemarcatedfor functional #urposes (see Fig. 2). These three partsare:

1. The caecal colon, which lies above or beyond theileo-caecal orifice; it is usually regarded as the commence-ment of the ascending colon, but a comparative studyshows that it represents the primitive caecum of the lowervertebrates (Fig. 2).

2. The caecum proper, which lies below or proximal tothe ileo-caecal orifice, and arises as a diverticulum fromthe caecal colon,

=

i6oo TM Enjurass 'IBfZDIDAL JOUR14AL j [DEC. 79 19112,6

Page 3: THE FUNCTIONAL NATURE OF THE CAECUM AND APPENDIX.*

,2*UNCTION 0 CAECUMI AND APPFNDIE. ITHE BTRIMSEI mErDICAL JOURNAL I6or

3. The apical or appendicular part, which is charac-terized by its narrow lumen and its thicker walls. Pro-fessor Berry'3 showed twelve years ago that it is linedby a mucous membrane laden with lymphoid tissue.The same three divisions are to be recognized in the

human ileo-caecal regrion-caecal colon, caecrum andappendix (see Fig 1). In the rat's eaccum there is a dis-tinct sphincteric ring (caecal sphincter) at the junction ofthe caecum and caecal colon. In the human caecum thissphinteric ring (caecal sphincter) is permanently unitedwith the ileo-caecal orifice, and forms the well-knownretinacular bands (Fig. 1). The action of the caecalsphincter can be studied in the rat (see Fig. 3). At oneminute (Fig. 3, A) the ileo-caecal orifice is closed and theeaccal sphincter dilated; at another this sphincter is in-action (Fig. 3, B), and the ileum opens directly into thecaecum; other states are represented in Fig, 3, C, D. Asyet we know nothing of the action of the retinacula of thehuman caecum, but from what we know of their action inotlher animals we may safely infer that during caecaldigestion they can regulate the passage of the colic andiliac Contents to and from the caecum.

CAECO-COLIC SPHINCTERIc TRACT.In mammals which have the caecumn well developed,

the first part of the colon-corresponding to the com-mencement of the ascending colon in man-is differentiatedfrom the eaccum below and the rest of the colon above bythe degree to which its circular musculature is contracted(see Figs. 1 and 2). I regard this part as serving as a

COLON

S:T,SeSPH: TRA,CT z<

IL:

CAMCUM CC:

A,

I L.

C.B. D

Fig. 3.-Figures illustrating four functional states of the ileo-caecal region of the bowel of rats, In A and c the ileo-caecalsphincter is closed; in B and D it is relaxed. -In A and B thesphincteric tract is closed; in c and D it is open. Variousconditions of the caeco-colic sphincter are shown in A, B, c, and D.

functional sphincter of the eaecal colon.- It is true thatnot one of the many experts who have watched thepassage of a bismuth-laden diet have been able to verifythe existence of this sphincteric tract. It must be remem-bered that it is hard to be certain of the movements inthe ileo-caecal region; the bony pelvis prevents us fromhaving a clear image, and the ascending colon awvavsbends downwards in front of the caeco-colic junction, sothat it is difficult to watch the passaae of bismuth-ladenchyme as it passes into the beginning of the colon. Inthe routine of abdominal operations the functional dif-ferentiation of the segments of the bowel is not observedowing to the paralytic effect of anaesthetics; in theroutine of the _ost-mortem room the functional states arenot seen because the putrefactive action of the caecal andcolic contents soon destroys the vital action of the muscularcoat -of the great bowel. The three reasons -which con-vince me I am right in differentiating a functionalsphincteric tract are:

1. Its presence in all mammals with well-developedcaeca. Comparative anatomists have failed to recognizeit because in some animals, such as the horse, the caecalcolon has become highly specialized and elongated, thusthrusting the sphincteric tract some distance away fromthe ileo-caecal orifice.

2. In young and healthy subjects, if formalin be injectedby the arterial system, before putrefaction has destroyedthe vitality of the musculature of the bowel, this tract willbe seen (see Fig. 1 and also Professor Symington's draw-ing in Quain's Anatomy, vol. iii, Part IV, p. 190, 1896).

3. It is impossible to decompress gas from the caecumof the living; the caecum always contains gas; that is anecessary correlative of the process of digestion going onwithin it. The gas is locked up in the caecum just as itis in the cardiac part of the stomach; it cannot be pressedinto the ileum because of the ileo-caecal sphincter; it

cannot be pressed along the ascending colon because ofthe sphincteric nature of the muscular wall' at theBcommencement of the ascending colon.

THE ILEO-COLIC REGION OF BIRDS.The ileo-colic region reaches its llighest specialization

in grasninivorous and vegetable feeding birds. A study ofthe avian mechanism helps us very materially to under-stand the working of the ileo-caecal tract in mammals.In Fig. 4 I have represented the termination of the ileum;and beginning of the colon of a young ostrich. Betweenthe ileum and colon there is a well-demarcated chambercorresponding to the caecal colon of mammals. It isseparated from the ileum by a sphincter (ileo-colic),-andfrom the colon by a caeco-colic sphincteric tract (Fig. 4).The two long caeca open from the caecal colon by acommon orifice, which is guarded by a strong caecalsphincter. It is easy to see that the caecal colon is thefeeder of the caeca; when it contracts, its contents, beingshut off by the sphincters at its iliac and colic ends, areforced into the caeca or onwards into the colon.

It appears, however, that the caecal colon has also aselective function. In the valuable report"4 by the com-mittee which recently inquired into grouse disease-Dr. Hammond Smith drew my attention to this sourceof information-a number of valuable observations arerecorded. The rough fibres and debris of heather shootsanld of moor fruit present in the chyme when it reachesthe colic caecum are passed on to the colon through thesphincteric tract, while the more fluid matter is pressed into

JonllM I 1;%SASPHFRI-P C)'ECUM

/ C.4VI ILEO-CgC: ORIFICEI CA!:CAL COLON

CZICAL ORIFICE

Fig. 4.-The ileo-caecal region of a young ostrich.

the cacculmi. Thus, the caeca, which together are twicethe length of the small bowel in the grouse, are filled withfluid, alkaline chyme from the ileum; in the caeca itsreaction becomes acid, probably from the bacterial diges-tive processes whicll take place in the caeca. The caecaare only partially and occasionally emptied, the caecalcontents differing completely in character from the usual" droppings'" of the grouse. Here, then, in its highestform we find the caecal colon acting as a selectivemechanism and the caeca set aside as separate chambersfor a special and apparently bacterial process of digestionand absorption.

EMPTYING OF THE CAECUM.In rodents the caecuin begins to pass into action as soon

as food enters the stomach, but at no time does thecaecum of any animal become completely emptied. It iswell knownv that in animals which die of starvation thecaecum and adjacent part of the ascending colon retaintheir normal contents, but the significance of this fact hasnot received the attention its importance merits. If thedigestive processes undergone by food in the caecum andgreat bowel, are due to bacterial action, then it is abso-lutely necessary that a caecal " batch " must be preservedto secure the continuation of bacterial digestion. Obser-vations made by Elliott 'and Barclay, Smith have animportant bearing on the mechanism which seciures theretention of the caecal batch. In carnivores, such as thecat, ferret, dog, stimulation of the sacral visceral nervescauses a constriction and emptying of the great gut fromthe caecum to the rectum. In vegetable-eating animals theeffect extends to only the distal part of the colon; theproximal part of the colon-corresponding to the caecum,ascending colon, and the hepatic part of the trans-verse colon of man-are outside the influence of thisemptying mechanism. Holzknecht and Dr. A. E.

D)EC. 7,. I9I12.]

-UKA(;TI

Page 4: THE FUNCTIONAL NATURE OF THE CAECUM AND APPENDIX.*

1602 yT=Bwno I FUNCTION OF CAE-CUM AND APPENDIX, [DEC. 7, 1912.,

Barclay15 have observed the peculiar expulsory move-ments in the spleBiic part of the transverse colon. Weknow that expulsory movements occur in the colon atand distal to the splenic flexure. We must infer thatthe preservation of part of the caecal and colic con-tents is essential for the digestive economy of vegetable-feeding mammals, and if we admit that the digestiveprocesses of the caecum and colon result from theaction of certain bacteria we see why Nature has placedthe proximal part of the colon outside the influence ofthe expulsory mechanism. In the proximal part of thecolon antip.ristaltic or retaining movements are observed,never the movements of complete expulsion. For thesame reason we can explain the absence of the greatbowel in all forms of fishes; the differentiation of thebowel into small and great intestine 'appears with theevolution of land-living, air-breattling vertebrates. Abacterial form of digestion, necessarily attended by theproduction of gas within the bowel, would upset thestability of gill-breathing, free-swimming forms. Thedigestive juices of fishes are notoriously strong. From thepoint of view of comparative anatomy, we must regardthe great bowel as a part which has been elaborately'modified and evolved, through the course of long ages, fora highly peculiar form of digestion.Some years ago I noted the following facts relating to

the digestive changes in the bowel of rats fed on oats.In the jejunum free starch was abundant; in the lowerpart of the ileum no starch reaction was obtained. It wasnot until the contents of the colon had reached the midpart of the transverse colon that the starch reaction becameagain manifest. These observations seem to indicatethat cellulose digestion had proceeded in the proximalpart of the colon, exposing fresh granules of starch whichwere absorbed in the middle and distal part of the colon.

THE ALLEGED VESTIGIAL NATURE OF THE APPENDIX.For many years the appendix vermiformis has been

regarded as one of the vestigial structures of man's body,an opinion which has prejudiced us against any realendeavour to discover its nature and function. Twelveyears ago Professor R. J. A. Berry, on the evidence hethen produced, came to the conclusion that the appendixshould be regarded as a specialized part-not a retrogradepart-of the caecum, and that is the conclusion whichevery one must reach who* makes an impartial survey ofour knowledge of the comparative anatomy and of the evo-lution of the human ileo-caecal region. At least in threeseparate orders of mammals the apical part of the caecumhas become specialized and demarcated from the rest oftlle caecum in the form of an appendix. It has neverbeen suggested that the appendix of the koala, wombat,rabbit, lemur, or of the anthropoid apes is a vestigialstructure. So like are the caeca and appendix of thegorilla and chimpanzee to the same structures in man,that I know of no character by which an expert could tellthe one from the other. In the anthropoids, as in man,the caecum and appendix undergo the same peculiarchanges in passing from infancy to adult age. We knowfrom their fossil remains that the great anthropoids are ofextreme geological antiquity; were their appendices in-jurious or vestigial structures, there has been ample timeto accomplish its complete suppression. Kelly andHurdon, in their well-known work,'6 cite the small calibreof the appendix as evidence of its retrograde character;it is more in keeping with what we know of animal struc-tures to regard the smallness of calibre as an adaptationto a peculiar but unknown function. We do not supposethe oesophagus to be a retrograde organ because it is narrowas compared to the stomach; we fortunately know thefunction of the gullet.

PROBABLE CAUSE OF DERANGEMENT.Every one who is acquainted with the more recent

developments of our knowledge of the surgery andpathology of the abdomen agrees that, in a large pro-portion of civilized men and women, the great bowel-especially the caecum and appendix-is abnormally liableto derangement and to disease, but as to the cause of thatcondition there is a diversity of opinion. No observationsof recent years throw more light on appendicitis thanthose which Dr. Weinberg'7 made on chimpanzees dyingin captivity. We have seen that the caecum and appendix

have the same form in the chimpanzee as in man. In tenout of sixty-one chimpanzees appendicitis was found atdeath. We have no reason to suppose that in its naturalhabitat this anthropoid is specially liable to appendicitis-the evidence is purely negative;- but as soon as the chim-panzee comes into captivity, and is placed on a human'diet and exposed to human contagion, it becomes subjectto a prevalent human disease. In the chimpanzee's casewe blame the diet. Metchnikoff and Dr. Barclay Smith,as far as man is concerned, regard the appendix as at fault.When we think of how the diet of highly civilized raceshas changed-in quality, quantity, and character-incomparatively recent times, one must marvel that ourorganization, which was evolved to deal with a moreprimitive and more precarious supply of food, has accom-modated itself to modern conditions so well as it has. Weknow that beyond the neolithic period, when cereals beganto be cultivated, some six thousand years ago, there liesa vast hinterland of rude human existence, when manmust have lived on the natural products of the country.With the discovery of fire and of the artificial preparationof food (we know that man had discovered the use of fire.before the end of the Pleistocene period) the task of thealimentary system must have been greatly altered. Thegreatest changes, however, are those of more recent cen-turies-the concentrated nature of food, its plentifulsupply, its highly artificial character. When we come torealize how slowly evolutionary processes have affectedman's body in past times, we can hardly expect ourinternal digestive system to adapt itself to the rapid pacedemanded by the ever-accumulating resources of civiliza-tion. The modern changes we see at work in our teethand jaws have set in since neolithic times; we have everyreason to suppose these are allied to, and contemporaneouswith, changes affecting the whole alimentary system.Thus an impartial survey of the evidence at present at

the disposal of the anatomists indicates very plainly thatwe cannot hope to prevent or cure the ailments to whichthe great bowel is liable so long as we regard it as a.hopelessly injurious or useless structure. On the otherhand, if we regard it as having all the anatomical appear-ances of a useful structure, our outlook becomes hopeful ifwe can only discover what its uses are. If we only knewhow to keep it suitably and profitably employed by alter-ing our diet to meet its requirements, it will, we haveevery reason to think, serve us and future generations justas well as it answered the digestive needs of primitive andsuccessful races in the past.

REFERENCES.1E. Barclay Smith, Proc. Camb. Phil, Soc., October 27th, 1902, vol.

xii. 2Elie Metchnikoff, The Nature of Mfan, English edition by P.Chalmers Mitchell, 1903. W. Arbuthnot Lane, Lancet, 1903, i, p. 153;1904, i, p. 19. 4R. Owen, Hunter's Essays and Observations, vol. i.p. l48. 5R. Owen, The Anatomy of Vertebrates, 1868, vol. iii, p. 426.6A. F. Hertz, BRiTISH MEDICAL JouRNAL, 1908, January 18th and 25th.7W. B. Cannon, TheMechanical Factors of Digestion, 1911. 8A. Keith,Proc. Anat. Soc. Great Britain and Ireland, November, 1903, 1p. 1-London Hospital Med. Gaz., November, 1903, p. 62. 9For an accountof the comparative anatomy of the caecum and appendix, see G. S.Huntingdon, The Anatomy of the Human Peritoneum and AbdominalCavity, London, 1903. i0 T. R. Elliott, Tourn. of Physiol.. 1904, vol. xxxi,p. 157. ILT. R. Elliott and E. Barclay Smith, Journ. of Physiol., 1904,vol. xxxi, p. 272. 12 Sir Wm. Macewen, BRITIsH MEDICAL JOURNAL,October 8th, 1904. 1R. J. A. Berry, J7ourn. Anat. and Pht,siol., 1900,vol. xxxv. p. 83, 1906, vol. xl, p. 274; Intercolonial Med. Tourn. ofAustralasia, June 20th, 1907. 14The Grouse in Health and Disease.Report of Committee of Inquiry, 1911. 15A. E. Barclay, Arch. RoentgenRays, April, 1912. 16 Howard A. Kelly and E. Hurdon, The Vermiform4Afppendix and Its Diseases, 1905. 17 Weinberg, Bull. de la Soc. de Path.,November, 1908, p. 556.

WITH the approval of King Edward's Hospital Fund forLondon and the Charity Commissioners, an amalgamationof the British Lying-in Hospital with the Home for Mothersand Babies, Woolwich, the President of which is H.R.H.,Princess Christian, has been provisionally arranged underthe title of the British Hospital for Mothers and Babies..The subtitle of the latter institution, " National TrainingSchool for District Midwives," will be retained, and thejoint hospital will occupy a new building, which is shortlynto be erected at Woolwich for the purpose. The authoritiesof the British Lying-in Hospital have been led to this stepby the rebuilding and alteration in character of the district,around Endell Street, which has greatly reduced thenumber of its patients, whilst the addition of maternitywards to most of the large hospitals has also influencedtheir decision. They feel that the usefulness of the insti-tution will be materially increased if it follows thepopulation to the outskirts of LJondon.